ArticlePDF Available

Abstract

Objective Fisheries provide countless benefits to human populations but face many threats ranging from climate change to overfishing. Despite these threats and an increase in fishing pressure globally, most stocks remain unassessed and data limited. An abundance of data‐limited assessment methods exists, but each has different data requirements, caveats, and limitations. Furthermore, developing informative model priors can be difficult when little is known about the stock, and uncertain model parameters could create misleading results about stock status. Our research illustrates an approach for rapidly creating robust initial assessments of unregulated and data‐limited fisheries without the need for additional data collection. Methods Our method uses stakeholder knowledge combined with a series of data‐limited tools to identify an appropriate stock assessment method, conduct an assessment, and examine how model uncertainty influences the results. Our approach was applied to the unregulated and data‐limited fishery for Crevalle Jack Caranx hippos in Florida. Result Results suggested a steady increase in exploitation and a decline in stock biomass over time, with the stock currently overfished and undergoing overfishing. These findings highlight a need for management action to prevent continued stock depletion. Conclusion Our approach can help to streamline the initial assessment and management process for unregulated and data‐limited stocks and serves as an additional tool for combating the many threats facing global fisheries.
Marine and Coastal Fisheries: Dynamics, Management, and Ecosystem Science. 2023;15:e10270.
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1 of 22
https://doi.org/10.1002/mcf2.10270
wileyonlinelibrary.com/journal/mcf2
Received: 18 November 2022
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Revised: 20 July 2023
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Accepted: 3 August 2023
DOI: 10.1002/mcf2.10270
ARTICLE
Rapid approach for assessing an unregulated fishery using a
series of data- limited tools
Carissa L.Gervasi1
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MandyKarnauskas2
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AdyanRios2
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Rolando O.Santos1
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W. RyanJames1
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Ryan J.Rezek3
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Jennifer S.Rehage1
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided
the original work is properly cited.
© 2023 The Authors. Marine and Coastal Fisheries published by Wiley Periodicals LLC on behalf of American Fisheries Society.
1Institute of Environment, Florida
International University, Miami,
Florida, USA
2National Oceanic and Atmospheric
Administration, Southeast Fisheries
Science Center, Key Biscayne, Florida,
USA
3Department of Marine Science,
Coastal Carolina University, Conway,
South Carolina, USA
Correspondence
Carissa L. Gervasi
Email: carissa.gervasi@noaa.gov
Funding information
National Science Foundation, Grant/
Award Number: HRD- 1547798 and
HRD- 2111661
Abstract
Objective: Fisheries provide countless benefits to human populations but face
many threats ranging from climate change to overfishing. Despite these threats
and an increase in fishing pressure globally, most stocks remain unassessed and
data limited. An abundance of data- limited assessment methods exists, but each
has different data requirements, caveats, and limitations. Furthermore, developing
informative model priors can be difficult when little is known about the stock, and
uncertain model parameters could create misleading results about stock status. Our
research illustrates an approach for rapidly creating robust initial assessments of un-
regulated and data- limited fisheries without the need for additional data collection.
Methods: Our method uses stakeholder knowledge combined with a series of
data- limited tools to identify an appropriate stock assessment method, conduct
an assessment, and examine how model uncertainty influences the results. Our
approach was applied to the unregulated and data- limited fishery for Crevalle
Jack Caranx hippos in Florida.
Result: Results suggested a steady increase in exploitation and a decline in stock
biomass over time, with the stock currently overfished and undergoing overfish-
ing. These findings highlight a need for management action to prevent continued
stock depletion.
Conclusion: Our approach can help to streamline the initial assessment and
management process for unregulated and data- limited stocks and serves as an
additional tool for combating the many threats facing global fisheries.
KEYWORDS
CMSY, Crevalle Jack, data limited, FishPath, local ecological knowledge, stock assessment
INTRODUCTION
Despite their importance, the status of many global fish-
eries remains unknown or poorly estimated due to a lack
of sufficient data or institutional capacity required to con-
duct traditional stock assessments (Cope et al. 2023). The
majority of global fisheries are lacking formal assessment,
and studies have estimated that these unassessed fisher-
ies may be in significantly worse condition than assessed
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GERVASI et al.
fisheries (Costello et al.2012; Blasco et al.2020; Hilborn
et al. 2020). Furthermore, due to increasing fishing pres-
sure and constraints on fisheries management programs,
the development of monitoring and assessment plans for all
harvested fish species is an unattainable goal (Harford and
Carruthers2017; Sagarese et al.2019). Although significant
progress has been made toward improving fisheries data col-
lection (e.g., Bryan et al.2016; Amoroso et al.2018; Rousseau
et al.2019), there will likely continue to be a need for alter-
native, data- limited approaches to stock assessment in the
future (Sagarese et al. 2019). This is particularly true for
areas like the southeastern United States, a highly biodiverse
region where fisheries are dominated by the recreational
sector (Shertzer et al.2019) and where over 75% of stocks are
considered data limited (i.e., lacking sufficient data to con-
duct traditional assessments; Berkson and Thorson 2015;
Newman et al.2015). There is an urgent need for (1) rapid
assessment and management action that can keep pace with
increasing fishing pressure and (2) methods that can iden-
tify unregulated and data- limited fisheries that are at risk of
overexploitation and depletion (Sun et al.2020).
Over the past few decades, numerous data- limited
assessment methods have emerged to tackle this issue
(Dowling et al.2015). Rather than relying on traditional
quantitative, model- based stock assessments, these meth-
ods estimate the status of fish stocks by using a range of
approaches from expert judgment to multiple indicator
models (Dowling et al. 2019). However, methods differ
greatly in their data requirements, caveats, and context,
making it difficult to determine which assessment method
is the best choice for a particular fishery. Blanket applica-
tion of generic models can lead to an inaccurate portrayal
of fishery status and trends, thereby hindering effective
management (Dowling et al. 2019). This is because using
generic methods without first assessing whether they are
suitable for the fishery of interest increases the likelihood
of violating model assumptions and overlooking biases or
other data quality issues. Fortunately, several decision sup-
port tools have been developed in recent years that aim to
assist fisheries scientists, managers, and stakeholders in de-
termining the appropriate methods for assessing and man-
aging a given fishery (McDonald et al.2018). One example
is the FishPath tool (www.fishp ath.org), a decision support
tool that was developed in 2016 and helps to guide users
through the selection of appropriate methods for monitor-
ing, assessment, and management of data- limited fisheries
(Dowling et al.2016). The FishPath online assessment tool
contains a repository of data requirements and assump-
tions for over 50 stock assessment methods, with a focus
on data- limited options (Fitzgerald et al. 2018; Dichmont
et al.2021). Users first characterize their fishery via a series
of multiple- choice questions concerning biological and
life history attributes, fishery operational characteristics,
data availability, socioeconomic factors, and governance
context. The answers to these questions are then used to
identify possible assessment and management options that
are best suited to the fishery. Using a standardized tool like
FishPath can provide consistency and objectivity to data-
limited fisheries management and has the potential to be-
come a key resource for the assessment and management
of unregulated species (Fitzgerald et al.2018).
In addition to the development of numerous alternative
approaches to traditional stock assessment, fisheries science
is increasingly using stakeholder local ecological knowledge
(LEK) to help identify conservation concerns (Silvano and
Valbo- Jørgensen2008; Gervasi et al.2022b), estimate trends
in stock status over time (Beaudreau and Levin2014; Kroloff
et al. 2019), improve fisheries models (Bélisle et al. 2018),
and fill in critical knowledge gaps about species biology and
ecology (Anadón et al.2010). Local ecological knowledge is
the in- depth knowledge of the local natural environment ob-
tained by individuals or groups of people through personal
observations, practical experience, and community dialog
(Anadón et al.2009). Research has shown that angler LEK
can complement biological data and provide new insights
(Silvano et al.2008; Cardoso da Silva et al.2020). For example,
Figus et al.(2017) showed that both fishermen and scientists
observed similar declines in the abundance and condition
of Atlantic Cod Gadus morhua in the eastern Baltic Sea,
Poland. In addition to this consensus, fisher LEK revealed
a potential driver of the decline that was at odds with the
findings of scientists, prompting additional avenues for re-
search. There are several examples of angler LEK being used
to directly inform fisheries management, including develop-
ing management options with a high probability of success
and compliance (Heyman and Granados- Dieseldorff2012),
understanding causes of disagreement with existing man-
agement measures and differing stakeholder preferences
(Hill et al.2010; Figus and Criddle2019), developing fishery
surveillance indicators that can be used to continually mon-
itor fisheries (Shephard et al.2021), and providing estimates
of model parameters used in stock assessments (Ainsworth
and Pitcher2005; Beaudreau and Levin 2014; Friedlander
et al.2015). Although these studies demonstrate clear bene-
fits to incorporating angler LEK into fisheries management,
Impact statement
This study presents a method for conducting
rapid, low- cost fish stock assessments that was
applied to the Crevalle Jack fishery in Florida.
Application of this method to unregulated fish
species can help managers better assess fish stocks
and conserve important fisheries.
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
there has been a lack of standardized protocols and methods
for doing so (Hind2015) and integration of LEK into bio-
logical assessments remains uncommon (Figus et al.2017).
The goal of this study was to develop an approach for
conducting rapid initial assessments (i.e., using only ex-
isting information sans additional data collection) of un-
regulated and data- limited fisheries that could be applied
to the Crevalle Jack Caranx hippos fishery in Florida. The
Crevalle Jack is a large marine species that is targeted by
both commercial and recreational anglers, but the fishery
in Florida is currently unregulated and data limited. Fur-
thermore, research has suggested that the population may
be in decline (Gervasi et al.2022b). Our approach used
angler LEK in conjunction with a series of data- limited
assessment tools to assess the current status of the Florida
Crevalle Jack stock, examine trends in stock status and ex-
ploitation over time, and develop initial management ref-
erence points, which are benchmarks that scientists and
managers use to set targets or limits on fishing effort and
to monitor the success of management strategies (Caddy
and Mahon1995). First, we used information gathered
from LEK and other sources to fill out the FishPath as-
sessment questionnaire and to choose a data- limited stock
assessment method that was suited to the fishery of inter-
est. Second, we conducted a stock assessment using the
chosen method, with LEK informing unknown model pa-
rameters and filling in data gaps. Finally, simple sensitiv-
ity analyses were run to test how uncertain or unknown
parameters (including those estimated by LEK) affected
the estimates of stock status (Figure1).
METHODS
Study species
The Crevalle Jack is a large pelagic fish species with a
native range spanning the east coasts of North America
and Central America (Smith- Vaniz and Carpenter 2007).
In Florida, the Crevalle Jack is a popular sport fish spe-
cies that is highly valued by recreational anglers for its
strength and speed (Gervasi et al.2022b). Crevalle Jack are
also captured in commercial fisheries (mainly as bycatch)
throughout the state but are unregulated and understud-
ied. The Florida Fish and Wildlife Conservation Commis-
sion makes management decisions for all fisheries within
state waters; such decisions include setting gear restric-
tions, size limits, and bag limits and instituting closed sea-
sons. State waters extend 4.83 km (3 mi) from shore on the
east coast of Florida and 16.09 km (10 mi) from shore on
Florida's west coast (Figure2). Several species (including
the Crevalle Jack) are listed as unregulated species in the
state, which means that they have no specific regulations
regarding gear restrictions, size limits, bag limits, or closed
seasons. Florida does, however, have a default limit of two
fish or 45.36 kg (100 lb) per person per day (whichever is
greater) for all unregulated species (Florida Fish and Wild-
life Conservation Commission [FWC]2021). Crevalle Jack
are found in a variety of habitats, including offshore reefs
(Smith- Vaniz and Carpenter 2007). Hence, they are also
captured in federal waters within the U.S. Exclusive Eco-
nomic Zone (Figure2). Federal fisheries in the region are
managed by the National Oceanic and Atmospheric Ad-
ministration (NOAA) Fisheries, the South Atlantic Fish-
ery Management Council, and the Gulf of Mexico Fishery
Management Council (NOAA– Fisheries 2021). Crevalle
Jack are not currently managed as a federal species, so
there are no restrictions on their harvest in federal waters.
Due to the Crevalle Jack's unregulated status, limited
research has been done in the region to assess the species'
life history (e.g., stock boundaries and migration patterns)
or fishery trends (e.g., trends in length and age compo-
sition; McBride and McKown2000; Gervasi et al. 2022b;
Jefferson et al.2022). Therefore, the Crevalle Jack fishery
in Florida can be considered data limited, a term that gen-
erally describes situations in which the data required to
support a fully integrated stock assessment model (includ-
ing catch time series, indices of abundance, length and
age composition, and life history parameters) are missing
(Cope et al. 2023). In the Florida Keys, recreational fish-
ing guides have observed a concerning decline in Crevalle
Jack catch rates, which is supported by available fisheries-
dependent data (Gervasi et al. 2022b). This decline
prompts a pressing need for assessment of the species in
Florida and possible future management action. The Flor-
ida Crevalle Jack fishery is therefore an ideal candidate for
applying the methodology outlined herein.
The FishPath tool
In this study, the FishPath assessment questionnaire was
filled out by the lead author for Florida Crevalle Jack by
using information compiled from various sources (Table1;
File S1 available in the Supplement separately online).
When possible, published literature and existing fisheries-
dependent data were used to answer the multiple- choice
questions. However, some questions could not be an-
swered without additional research or data collection. In
these instances, LEK was used to fill in the knowledge
gaps. Fortunately, LEK data concerning the Crevalle Jack
population in south Florida were already available from
interviews conducted in 2019 with expert recreational fish-
ing guides in the Florida Keys (Gervasi et al.2022b). Most
of the remaining FishPath questions could be answered
using this existing LEK data set. Interview methods are
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GERVASI et al.
described in depth by Gervasi et al.(2022b). Briefly, key
informant interviews were conducted with 18 veteran
charter- for- hire captains in the Florida Keys, where a de-
cline in Crevalle Jack catches had been observed. Captains
were asked a series of open- ended questions to guide the
conversations: (1) “What is your general background and
experience fishing and guiding?”; (2) “What do you know
about Crevalle Jack?”; (3) “Have you noticed any changes
in Crevalle Jack fishing over time?”; and (4) “Is fishing for
Crevalle Jack important to you?” More specific follow- up
questions were asked as needed, with the goal of capturing
perceptions about the Crevalle Jack fishery and stock sta-
tus as well as gaining an understanding of how and why
stakeholders interact with the species (FigureS1 available
in the Supplement separately online). Common percep-
tions of the fish population and fishery among anglers
were summarized and used in the current study to fill
out the FishPath questionnaire. All protocols for human
subject research were approved by Florida International
University's (FIU) Institutional Review Board, and all par-
ticipants gave consent before being interviewed. Any re-
maining FishPath questions were answered by consulting
FIGURE Framework presented in this paper for conducting rapid initial assessments of unregulated and data- limited fisheries using
a three- prong approach, with angler local ecological knowledge (LEK) permeating each step. First, data from LEK and other sources are
used to fill out the FishPath assessment questionnaire and choose a data- limited stock assessment method suited to the fishery of interest.
Second, a stock assessment is conducted for the species of interest by using the chosen method, with LEK informing unknown model
parameters. Finally, simple sensitivity analyses are run to test how uncertain or unknown parameters (including those estimated by LEK)
affect estimates of stock status. Highly influential parameters highlight critical future research needs.
19425120, 2023, 5, Downloaded from https://afspubs.onlinelibrary.wiley.com/doi/10.1002/mcf2.10270, Wiley Online Library on [05/11/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
with the previously interviewed anglers and asking them
to address the specific question.
The FishPath assessment questionnaire includes five cat-
egories of questions concerning the biology and life history
of the species, data availability, governance, management,
and operational characteristics. All available fisheries-
dependent and fisheries- independent surveys that operate
in Florida and regularly encounter Crevalle Jack were com-
piled to answer questions about data availability. Previous
literature on the Crevalle Jack was used to inform ques-
tions about biology and life history (Smith- Vaniz and Car-
penter2007; Caiafa et al.2011; Alfaro- Martínez et al.2016;
Jefferson et al.2022). Common perceptions of the Crevalle
Jack fishery from LEK interviews were summarized to gen-
erate a LEK data set for use in the FishPath assessment and
the resulting selected model. The LEK data included infor-
mation on relative stock status and the nature of fishery op-
erations, including targeting, species uses, and fishing areas
(Table1; FileS1).
The FishPath assessment tool does not rank the possi-
ble assessment methods, but it does filter out any methods
for which the minimum data requirements or criteria are
not met based on the questionnaire responses (Dowling
et al.2016). The tool also displays “traffic light” caveats that
highlight each possible assessment method's major assump-
tions and data requirements as they relate to the fishery of
interest. Caveats that are red are important assumptions that
might not be met according to the questionnaire responses,
so those methods should be used with extreme caution. All
options with one or more red caveats were eliminated. Fish-
Path also ranks each method by assessment tier (i.e., model
complexity), with tiers ranging from simple, extremely data-
limited methods (pre- assessment) to robust methods that
require additional data (high tier). Options were sorted by
assessment tier, and the highest tier options that remained
after the elimination of options with red caveats were con-
sidered the best options for assessment of the Florida Cre-
valle Jack fishery (FileS2).
FIGURE Map of the study area in Florida, highlighting state water boundaries (4.83 km [3 mi] from shore on the east coast; 16.09 km
[10 mi] from shore on the west coast) and the U.S. federal Exclusive Economic Zone (EEZ). The inset map highlights the study area in the
southeastern United States. (State boundary shapefile was downloaded from FWC- Fish and Wildlife Research Institute2007; federal EEZ
shapefile was downloaded from Flanders Marine Institute2019).
19425120, 2023, 5, Downloaded from https://afspubs.onlinelibrary.wiley.com/doi/10.1002/mcf2.10270, Wiley Online Library on [05/11/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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GERVASI et al.
Model inputs
Model priors
All methods pertaining to model inputs were contingent
upon the results of using the FishPath tool. We chose
the highest tier type of assessment recommended: the
CMSY– BSM model (CMSY = catch– maximum sustainable
yield [MSY]; BSM = Bayesian state- space implementation of
the Schaefer surplus production model; Froese et al.2017,
2019). Details of this selection are presented in the Results
(Assessment model selection). The CMSY– BSM model re-
quires a time series of total fishery removals (hereafter re-
ferred to as “catch”); priors for resilience (defined as species
TABLE Available data and information about Crevalle Jack that were compiled and used to fill out the FishPath assessment
questionnaire. FL, fork length; FWC, Fish and Wildlife Conservation Commission; GOM, Gulf of Mexico; LEK, local ecological knowledge;
MRIP, Marine Recreational Information Program; NOAA, National Oceanic and Atmospheric Administration.
Data category Variable Estimate Source
Biological and life
history attributes
Minimum size at maturity ~40 cm FL Caiafa et al.2011 (Colombia)
Maximum size ~100 cm FL Jefferson et al.2022 (GOM)
Longevity 20 years Jefferson et al.2022 (GOM)
Size at 50% maturity 62.6 cm FL (males); 66.2 cm FL (females) Caiafa et al.2011 (Colombia)
Length– weight
log10(weight)=−16.47 +2.79
log10(FL)
Jefferson et al.2022 (GOM)
Growth rate (females)
l
t
(F)
=903.04
[
1e
0.39(t0.73)]
Jefferson et al.2022 (GOM)
Growth rate (males)
lt(M)
=887.16
[
1e
0.39
(
t
0.73
)
]
Jefferson et al.2022 (GOM)
Length– fecundity Relationship not significant Alfaro- Martínez et al.2016
(Colombia)
Natural mortality rate 1.12 year−1 Caiafa et al.2011 (Colombia)
Fishing mortality rate 0.63 year−1 Caiafa et al.2011 (Colombia)
Stock boundaries Single Florida stock Preliminary acoustic telemetry data
(C. L. Gervasi, unpublished
data)
Habitat use Variable Smith- Vaniz and Carpenter2007;
fishing guide LEK (Gervasi
et al.2022b)
Available data Commercial landings Available for the entire state of Florida
from 1950 to 2021
Florida FWC
Commercial effort Number of landings receipts can be a proxy Florida FWC
Commercial size composition Size/age composition data are not available N/A
Recreational landings Available for the entire state of Florida
from 1981 to 2021
NOAA MRIP dockside survey
Recreational effort Estimated via household mail survey NOAA Fishing Effort Survey
Recreational size composition Subset of size data available for landed fish NOAA MRIP dockside survey
Fisheries independent No suitable surveys in the region N/A
Fishery attributes and
management
Stock status High in the 1980s; gradual decline since Fishing guide LEK (Gervasi
et al.2022b)
Commercial gear used Cast net, hook and line, gill net = 67% Commercial data
Recreational gear used Hook and line = 99% Recreational data
Discards >80% of recreational catch discarded annually Recreational data
Discard mortality rate 10% Fishing guide LEK (guides were
consulted specifically for this
study)
Targeting Opportunistic; some targeting Fishing guide LEK (Gervasi
et al.2022b)
Management Unregulated Florida FWC
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
productivity or resilience to fishing); priors for biomass B
relative to carrying capacity k (i.e., B/k) at the beginning,
middle, and end of the catch time series; and an optional
biomass time series. For Crevalle Jack, prior estimates for
the maximum intrinsic rate of population increase (r) were
extracted from the “Estimates based on models” section of
FishBase using a species resilience category of “medium”
(Smith- Vaniz et al. 1990; Froese and Pauly2021). A prior
range for k was derived by the model from the maximum
catch. The available time series of recreational fishing effort
used to develop an index of Crevalle Jack abundance did not
cover the entire time series of fishing effort. Therefore, fish-
ing guide LEK was used to determine prior range categories
for B/k at the beginning, middle, and end of the time series
based on guide estimates of how population abundance has
changed over time. The default B/k ranges corresponding to
these categories from Froese et al.(2017) were used in the
model (Table2).
Catch time series
The following formula was used to create a time series of
total fishery removals (catch time series) for Crevalle Jack
in Florida from 1950 to 2021:
where t is year and “discard mortality is an estimated
discard mortality rate (Figure3). Discard mortality occurs
when fish are caught and released alive but die after release
due to injuries suffered from the angling encounter or due
to an increased susceptibility to predation (Rudershausen
et al. 2007). The discard mortality rate is defined as the
proportion of individuals that suffer from discard mortal-
ity and therefore can be considered a component of fish-
ery removals. Previously interviewed fishing guides were
asked to estimate a discard mortality rate. The average of
their responses was used as the base discard mortality rate
in this study. Preliminary acoustic telemetry research has
revealed population connectivity of Crevalle Jack through-
out the state of Florida (C. L. Gervasi, unpublished data).
Hence, we assumed that state- level boundaries provided a
reasonable approximation of the stock unit. All catch data
were thus collected for the entire state. Commercial land-
ings data were obtained from the National Marine Fisheries
Service's Accumulated Landings System (NOAA 2021a).
Landings data for Crevalle Jack were downloaded for all
of Florida from the beginning of the time series (1950) to
the last available year (2021). Crevalle Jack recreational
landings (fish that were brought back to shore) and dis-
cards (fish that were caught and released either dead or
alive) for the state of Florida were downloaded from
the NOAA Fisheries' Marine Recreational Information
Program (MRIP) online query tool for the period of record
from 1981 to 2021 (NOAA 2021b). Additional details on
how commercial and recreational landings data were ob-
tained and how the discard mortality rate was calculated
are available in FileS3.
Abundance time series
No fisheries- independent surveys are operating in the
region that regularly encounter adult Crevalle Jack, but
relative abundance trends can be inferred from catch- per-
unit- effort (CPUE) data (Maunder and Punt 2004). For
the purposes of this study, we used the MRIP CPUE data
subset for Florida to create an index of Crevalle Jack abun-
dance for the entire state. Numerous factors besides stock
abundance can influence fishery catch rates (e.g., spatial,
temporal, and environmental variability), so we stand-
ardized the CPUE data for Crevalle Jack in Florida using
generalized linear models (GLMs; Matthews 2014). Spe-
cifically, a delta- lognormal GLM approach (Lo et al.1992)
was applied, with the following categorical factors in-
cluded in the model: year (1991– 2021), season (spring:
March, April, and May; summer: June, July, and August;
fall: September, October, and November; winter: Decem-
ber, January, and February), fishing mode (shore, charter,
or private), and day (weekday or weekend). Additional de-
tails on standardization methods can be found in Gervasi
et al. (2022b). Filtered and cleaned MRIP data included
240,712 trips from 1991 to 2021 (31 years). Of these trips,
Crevalle Jack were caught during 38,825 trips (16%). Based
on model selection via backward stepwise regression and
deviance tables, the final model for the proportion posi-
tive GLM included year and season as fixed factors, and
the final model for the positive trip GLM included year,
season, and fishing mode as fixed factors (TablesS1 and
S2 available in the Supplement separately online).
Sensitivity analyses
To examine the sensitivity of our Crevalle Jack CMSY–
BSM model to unknown or poorly estimated parameters,
we conducted a series of sensitivity analyses and compared
stock status as well as biomass and exploitation trends to
those generated from our base model (Table2). We ex-
plored seven different scenarios that tested the model sen-
sitivity to potential uncertainty by varying the estimated
discard mortality rate instead of using the LEK- derived
value (analysis 1); ignoring the CPUE data and using only
the CMSY model without the surplus production model
(1)
Commercial landings
t+
recreational landings
t
+
(
recreational discards
t
×discard mortality
)
,
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GERVASI et al.
TABLE Model inputs and parameters for the base Crevalle Jack stock assessment model and for the seven sensitivity analyses. B, biomass; CMSY, catch–maximum sustainable yield;
CPUE, catch per unit effort; ENP, Everglades National Park; k, carrying capacity; LEK, local ecological knowledge; MRIP, Marine Recreational Information Program; PSE, proportional
standard error; r, intrinsic population growth rate.
Variable Base model
Sensitivity
analysis 1
Sensitivity
analysis 2
Sensitivity
analysis 3
Sensitivity
analysis 4
Sensitivity
analysis 5
Sensitivity
analysis 6
Sensitivity
analysis 7
Description of sensitivity
analysis
Varying the
discard mortality
rate
Excluding CPUE
data (CMSY
model only)
Excluding
historical catch
Excluding high-
PSE points in
MRIP data
Using
uninformed
priors versus
LEK priors
Using an
alternative
abundance data
set
Accounting for
effort creep
Catch start year 1950 1950 1950 1981 1950 1950 1950 1950
Catch end year 2021 2021 2021 2021 2021 2021 2020 2021
Abundance time series MRIP MRIP N/A MRIP MRIP MRIP ENP MRIP
Resilience categoryaMedium Medium Medium Medium Medium Medium Medium Medium
Prior ranges for ra0.35– 0.80 0.35– 0.80 0.35– 0.80 0.35– 0.80 0.35– 0.80 0.35– 0.80 0.35– 0.80 0.35– 0.80
Relative biomass category
(beginning)b
Nearly unexploited Nearly
unexploited
Nearly unexploited Low depletion Nearly
unexploited
Uninformed Nearly
unexploited
Nearly
unexploited
Prior range for B/k
(beginning)b
0.75– 1.00 0.75– 1.00 0.75– 1.00 0.4– 0.8 0.75– 1.00 0.01– 1.00 0.75– 1.00 0.75– 1.00
Relative biomass category
(middle)b
Low depletion Low depletion Low depletion Low depletion Low depletion Uninformed Low depletion Low depletion
Prior range for B/k
(middle)b
0.4– 0.8 0.4– 0.8 0.4– 0.8 0.4– 0.8 0.4– 0.8 N/A 0.4– 0.8 0.4– 0.8
Relative biomass category
(end)b
Medium depletion Medium depletion Medium depletion Medium
depletion
Medium
depletion
Uninformed Medium depletion Medium
depletion
Prior range for B/k (end)b0.2– 0.6 0.2– 0.6 0.2– 0.6 0.2– 0.6 0.2– 0.6 0.01– 1.00 0.2– 0.6 0.2– 0.6
Effort creep (%) 0 0 0 0 0 0 0 2
Discard mortality rateb0.1 Range from 0.0
to 0.5
0.1 0.1 0.1 0.1 0.1 0.1
Note: Changes to model inputs and parameters from the base model for each sensitivity analysis are shown in bold.
aDerived from FishBase (Froese and Pauly2021).
bValues were derived from angler LEK except in sensitivity analysis 1 (discard mortality not informed by LEK) and sensitivity analysis 5 (priors not informed by LEK).
19425120, 2023, 5, Downloaded from https://afspubs.onlinelibrary.wiley.com/doi/10.1002/mcf2.10270, Wiley Online Library on [05/11/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
(BSM; analysis 2); subsetting the catch time series to begin
at 1981 (the beginning of the MRIP time series), thereby
ignoring the estimated historic recreational catch and his-
toric commercial catch (analysis 3); removing potential
outlier data points from the MRIP time series with high
proportional standard error (PSE; >50%) and replacing
them with interpolated values based on adjacent years
(analysis 4); using uninformed biomass priors instead of
the LEK- derived priors (analysis 5); using an alternative
standardized CPUE time series (developed by Gervasi
et al.2022b) based on the Everglades National Park (ENP)
creel survey (Osborne et al.2006), updated to include data
from 2020, as an index of abundance (analysis 6); and, fi-
nally, incorporating effort creep (2% annual increase in
catchability) into the model based on stock assessments
for other fish species in the Gulf of Mexico (Thorson and
Berkson2010; analysis 7). All analyses were conducted in
R version 4.2.3 (R Core Team 2023).
RESULTS
Assessment model selection
After eliminating FishPath assessment options with red
caveat outputs and sorting by assessment tier (highest to
lowest), two options with the high- tier designation re-
mained (File S2). These options were production models
(e.g., Schaefer, Fox, and Pella– Tomlinson models; Hil-
born and Walters1992) and the qR (catchability– recruits)
method (McGarvey et al. 1997). Production models re-
quire a continuous time series of fishery removals, and the
two major parameters in the models are r and k, which are
used to estimate MSY. Production models additionally re-
quire at least one index of abundance. The qR method uses
time series of catch by weight and in numbers, an estimate
of natural mortality (M), and an average of weight at age
to estimate biomass, catchability, exploitation rate, and
FIGURE (A) Time series of Crevalle Jack catch (total fishery removals; thousands of metric tons) used in the initial CMSY– BSM
model (defined in Methods) and (B) breakdown of fishery removals by fleet. In panels A and B, an estimated discard mortality rate of 10%
was applied to the recreational live releases to obtain an estimate of recreational dead discards. (C) The Marine Recreational Information
Program (MRIP) standardized abundance index, which was used in the base model and sensitivity analyses 1, 3, 4, 5, and 7 as the biomass
time series, is shown. (D) The Everglades National Park (ENP) standardized abundance index, which was used in sensitivity analysis 6 as the
biomass time series, is presented. Dashed lines in panels C and D are 95% confidence intervals.
Year
All fishery removals (1000 metric tons)
1950 1957 1964 1971 1978 1985 1992 1999 2006 2013 2020
0
1
2
3
4
5
6
7(A)
Year
All fishery removals (1000 metric tons)
1950 1957 1964 1971 1978 1985 1992 1999 2006 2013 2020
0
1
2
3
4
5
6
7
Recreational catch
Commercial catch
Recreational dead discards
(B)
Year
MRIP standardized abundance index
1990 1993 1996 1999 2002 2005 2008 2011 2014 2017 2020
0.0
0.5
1.0
1.5
2.0
2.5 (C)
Year
ENP standardized abundance index
1980 1984 1988 1992 1996 2000 2004 2008 2012 2016 2020
0.0
0.5
1.0
1.5
2.0
2.5 (D)
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GERVASI et al.
yearly recruitment. We thoroughly reviewed both meth-
ods in the scientific literature to select the best option for
assessment of the Crevalle Jack fishery. The qR method
was eliminated as an option because catch by numbers is
not recorded for Crevalle Jack in the commercial fishery
and would have had to be estimated. Furthermore, M for
Crevalle Jack in Florida is currently unknown and can-
not be estimated with any certainty (available estimates
are from areas outside the United States). The production
model required fewer inputs of uncertain parameters.
We chose to apply the CMSY– BSM method created
by Froese et al. (2017) and further updated by Froese
et al.(2019). This method includes both the BSM and the
CMSY model, which is similar to a production model but
does not require a time series of abundance. The CMSY–
BSM model was chosen because it estimates biomass, ex-
ploitation rate, MSY, and related fisheries reference points,
with the only data requirements being catch and productiv-
ity (Froese et al.2017). An extensive time series of fishery
removals in Florida is available for Crevalle Jack, and pro-
ductivity information is available from previous research.
The CMSY model is an updated version of the catch– MSY
method originally proposed by Martell and Froese(2013),
which reviews of data- limited assessment methods have
found to be a promising approach (International Council
for the Exploration of the Sea2014; Rosenberg et al.2014).
The predictions of the CMSYBSM method have been val-
idated against 48 simulated stocks and evaluated against
159 fully or partially assessed real stocks, and estimates
of r, k, and MSY were not significantly different from the
actual values for 90% of simulated stocks and 76% of real
stocks (Froese et al. 2017). Furthermore, a detailed user
manual and R code (last updated in 2019) are available for
download (Froese et al.2017), making the method easily
accessible and reproducible. The updated version of the
model (CMSY+ and BSM; Froese et al.2019) was used in
this study, but for simplicity we refer to it as the “CMSY–
BSM model” hereafter.
Base model run
Our base stock assessment model run for Crevalle Jack
revealed a gradual increase in exploitation and a corre-
sponding gradual decline in stock size from 1950 to 2021
(Figure4). Total catch was below MSY (3155 metric tons
year−1) from 1950 to 1988 and fluctuated around MSY for
the remaining years, with catch being above MSY for 19
of the 33 years from 1989 to 2021. Several definitions for
the terms “overfished” and “overfishing” exist in the lit-
erature, but generally a stock is considered “overfished”
if B is below BMSY by some degree and to be undergo-
ing “overfishing” if fishing mortality F is above FMSY by
some degree (Froese and Proelss 2012, 2013; Langseth
et al.2019; Hilborn2020). For the purposes of this study,
we referred to the Crevalle Jack stock as overfished when-
ever model- estimated B was below BMSY (B/BMSY < 1) and
as undergoing overfishing whenever model- estimated F
was above FMSY (F/FMSY > 1). These definitions do not ac-
count for fluctuations around the thresholds due to inher-
ent variability. A stock that is managed at MSY could be
expected to fluctuate around BMSY. However, the Crevalle
Jack stock is unmanaged and stock size has continually
declined, suggesting that the stock is not being sustain-
ably harvested. According to the base model, F was above
FMSY for 14 of the 22 years since 2000, revealing that over-
fishing has been occurring regularly since 2000. Biomass
was above BMSY from 1950 to 2002 but was below BMSY for
every year from 2003 to 2011 and from 2017 to 2021, with
B being the lowest in 2019. It appears that high levels of
catch above MSY starting in 1989 led to overfishing be-
ginning in 2000 and led to the stock becoming overfished
starting in 2003. According to the assessment, the current
status of the stock is overfished and undergoing overfish-
ing: the estimated F2021/FMSY was 1.12, and the estimated
B2021/BMSY was 0.88 (Table3).
Model sensitivity runs
Our first set of sensitivity runs examined the impact of se-
lecting various discard mortality rates for the recreational
fishery by running a series of models with recreational
discard mortality ranging from 0% to 50% in 10% incre-
ments (Table2; Figure5). Discard mortality rates above
50% were not considered because they were deemed
highly unrealistic by fishing guides (who were asked spe-
cifically about discard mortality for this study). For all
model runs, exploitation in 2021 (F2021/FMSY) was similar,
ranging from 1.1 to 1.3. Stock size in 2021 (B2021/BMSY)
generally increased with an increase in discard mortality,
ranging from 0.81 at 0% mortality to 0.95 at 50% mortality
(Table3; Figure5B). Regardless of the discard mortality
rate used, the models revealed the same trend of gradually
increasing exploitation and gradually decreasing stock
size over time. The status of the stock in 2021 was over-
fished and undergoing overfishing regardless of the mor-
tality rate used. The choice of discard mortality rate had
little effect on the estimate of r in the model but greatly
affected the estimate of k, with k increasing linearly as the
discard mortality rate increased (Figure5C,D). Since k di-
rectly affects the MSY estimate, the estimated MSY also
increased linearly as the discard mortality rate increased.
At a discard mortality rate of 0%, estimated MSY was 2320
metric tons; at a discard mortality rate of 50%, estimated
MSY was 6790 metric tons (Table3).
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11 of 22
ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
The remaining sensitivity analyses (2– 7) examined the
effects of excluding the abundance time series (using the
CMSY model only), excluding historical catch data, ex-
cluding high- PSE data points from the MRIP data, using
uniformed priors versus LEK priors, using an alternative
abundance data set, and accounting for an effort creep
of 2% per year (Table2). Except for sensitivity analy-
ses 5 (uninformed priors) and 6 (alternative abundance
data set), each of the sensitivity model runs revealed the
same pattern of gradually decreasing stock size over time
(Figure6A). Furthermore, estimated B2021 was below BMSY
for all models and estimated F2021 was above FMSY for all
models except in sensitivity analyses 2 and 5 (Table3;
Figure6B). For all model runs, B was below BMSY and F
was above FMSY for at least 4 years of the 71- year time se-
ries (FiguresS3– S8).
Excluding CPUE data (analysis 2) and using unin-
formed biomass priors (analysis 5) led to the most opti-
mistic depictions of current stock status. For analysis 2,
excluding an index of abundance led to a trajectory of
stock status over time similar to that from the base model,
with stock size gradually declining while exploitation
gradually increased over time (FigureS3). For analysis 5,
starting and ending biomass priors (1950 and 2021) were
set to a wide range (0.01– 1.00), which told the model that
we had no information about stock status at the beginning
or the end of the time series (Froese et al.2019). The inter-
mediate biomass level was set to “NA,” which allowed the
model to estimate it from maximum or minimum catch
according to some simple rules (Froese et al.2017). This
version of the model showed a trajectory of gradually in-
creasing exploitation over time, which was the same as the
FIGURE Summary of information relevant for management of Florida Crevalle Jack from the base CMSY– BSM model (defined
in Methods): (A) catches (total fishery removals; thousands of metric tons per year) relative to maximum sustainable yield (MSY; dashed
line); (B) development of predicted relative total biomass (B/BMSY); (C) relative exploitation (fishing mortality F/FMSY); and (D) trajectory of
relative stock size (B/BMSY) as a function of fishing pressure (F/FMSY). Gray shading in panels A– C denotes 95% confidence limits for MSY,
relative biomass, and relative exploitation, respectively. The oval shape around the assessment of the final year triangle indicates uncertainty
(yellow = 50% confidence interval [CI]; gray = 80% CI; dark gray = 95% CI).
1950 1960 1970 1980 1990 2000 2010 2020
02468
Catch (total fishery removals)
Catch (1000 metric tons/year)
MSY
1950 1960 1970 1980 1990 2000 2010 2020
0.00.5 1.01.5 2.0
Stock size
B/BMSY
1950 1960 1970 1980 1990 2000 2010 2020
0.00.5 1.01.5 2.0
Exploitation
F/FMSY
0.00.5 1.01.5 2.0
0.00.5 1.01.5 2.0
Stock status
B/BMSY
F/FMSY
1950
1989
2021
50% CI
80% CI
95% CI
(A)
(C) (D)
(B)
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12 of 22
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GERVASI et al.
base model's trajectory, but it estimated that stock size in
1950 was below BMSY, rapidly increased to high levels in
the 1960s, and then gradually declined (FigureS6). The
most important difference between these models and the
base model was that excluding CPUE data and using un-
informed priors painted a more optimistic picture of stock
status, with the stock in 2021 still just below BMSY but also
below FMSY, suggesting that overfishing is not currently
occurring (Table3; Figure6B). Additionally, exploitation
was above FMSY for only a handful of years during the 71-
year time series.
The most pessimistic models were sensitivity analy-
ses 4 (excluding high- PSE data points) and 7 (including
2% effort creep). Two data points had PSEs above 50%:
1986 and 2009. The catch estimate for 1986 (3107 metric
tons) was similar to the average of the time series. How-
ever, the catch estimate for 2009 (7116 metric tons) was
anomalously high compared to the rest of the time series,
whereas the mean catch before 2009 was 2078 metric tons
(Figure3B). Removing the 2009 data point and replacing
it with an interpolated value brought the total catch for
2009 down to 2787 metric tons. The decrease in total catch
for 2009 had a negative effect on r, with estimated r de-
creasing from 0.55 to 0.49. This resulted in a lower esti-
mated MSY and FMSY and a more pessimistic stock status,
with the stock being more severely overfished and under-
going more severe overfishing in 2021 (Table3; Figure6).
Based on this model, the Crevalle Jack stock was under-
going overfishing for 16 years of the 71- year time series
(FigureS5). Effort creep is defined as some change in
catchability or nominal effort in a fishery over time due to
technological advancements (Palomares and Pauly2019),
such as major improvements in gear design, fish- finding
devices, or vessel capabilities, all of which increase effi-
ciency and therefore impact fishing mortality. The CMSY
BSM model allows the user to specify a linear annual
increase in catchability, which results in a decrease in
the CPUE index considered by the model. For this sen-
sitivity analysis, a 2% linear increase in catchability was
applied to the MRIP standardized abundance index based
on previous stock assessments in the region (Thorson and
Berkson2010). Although effort creep did not impact the
trajectory of stock status and exploitation over time, it led
to a much steeper decline in stock status since the early
2000s than the base model (FigureS8). Furthermore, es-
timated B was below BMSY for every year since 2003, with
B/BMSY almost as low as 0.5 in 2021, suggestive of a se-
verely overfished stock.
Using an alternative abundance time series (analysis
6) had little effect on estimated management reference
points. However, the ENP time series went back farther
in time than the MRIP time series, and the trajectory of
stock status over time differed slightly between the two
TABLE Estimated management reference points from the base Crevalle Jack stock assessment model and from the seven sensitivity analyses. B2021, biomass in 2021; BMSY, biomass at
maximum sustainable yield (MSY); CMSY, catch– MSY; CPUE, catch per unit effort; ENP, Everglades National Park; F2021, fishing mortality in 2021; FMSY, fishing mortality at MSY; k, carrying
capacity; LEK, local ecological knowledge; MRIP, Marine Recreational Information Program; PSE, proportional standard error; r, intrinsic population growth rate.
Variable
Base
model
Sensitivity
analysis 1
Sensitivity
analysis 2
Sensitivity
analysis 3
Sensitivity
analysis 4
Sensitivity
analysis 5
Sensitivity
analysis 6
Sensitivity
analysis 7
Description of
sensitivity analysis
Varying the
discard mortality
rate
Excluding CPUE data
(CMSY model only)
Excluding
historical
catch
Excluding high- PSE
points in MRIP data
Using uninformed
priors versus LEK
priors
Using an alternative
abundance data set
Accounting for effort
creep
r (year−1) 0.55 0.50– 0.55 0.55 0.55 0.49 0.46 0.50 0.45
k (1000 metric tons) 22.9 17.8– 54.2 22.9 23.4 24.3 29.0 25.1 27.8
MSY (1000 metric
tons)
3.15 2.32– 6.79 3.44 3.21 2.96 3.31 3.14 3.11
FMSY (year −1) 0.28 0.25– 0.28 0.31 0.28 0.24 0.23 0.25 1.81
BMSY (1000 metric
tons)
11.4 8.89– 27.1 11.3 11.7 12.2 14.5 12.6 13.9
F2021/FMSY 1.12 1.10– 1.13 0.93 1.10 1.32 0.95 1.04 1.81
B2021/BMSY 0.88 0.81– 0.95 0.99 0.88 0.79 0.99 0.87 0.55
F2021 (year−1) 0.31 0.28– 0.31 0.28 0.30 0.32 0.22 0.26 0.41
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
models. With the ENP time series as an index of abun-
dance, stock size declined rapidly from 1970 to 1985 be-
fore increasing back to historic levels and then gradually
declining from 1990 to 2020 in the same fashion as the
base model (FigureS7). For the years in which the two
abundance time series overlapped (1991– 2020), model re-
sults were very similar.
DISCUSSION
In this paper, we demonstrated how a variety of data-
limited tools, when used in combination, can aid in de-
veloping rapid yet robust assessments for data- limited,
unregulated fisheries, thus providing a basis for initial
management. Our approach took advantage of LEK
to inform both model selection and analysis. We used
LEK and other existing data sources to fill out the Fish-
Path assessment questionnaire, which is a currently
underutilized program that provides a transparent,
standardized approach for selecting an appropriate
stock assessment model. Local ecological knowledge
was then used again to parameterize the chosen model
when parameter estimates were unavailable from previ-
ous literature, which is the case for many data- limited
fisheries. Finally, by identifying unknown and uncertain
parameters and running sensitivity analyses to test their
effects on estimates of stock status, we developed some
clear goals and priorities for future research, which will
help to ensure that funding and effort are invested in the
greatest needs. The results of applying our framework
to assessing stock status of the Crevalle Jack in Florida
suggested that B has been below BMSY for 14 of the past
19 years and that the stock is currently undergoing over-
fishing (with F slightly above FMSY). Any increase in
fishing pressure will likely lead to a continued decline
in stock size. Fishing guides in the Florida Keys have
observed a gradual decline in Crevalle Jack catch rates
FIGURE Results of the sensitivity analysis examining recreational discard mortality (sensitivity analysis 1): (A) time series of catch
(thousands of metric tons) with discard mortality set at 0.0, 0.1, 0.2, 0.3, 0.4, or 0.5; (B) time series of exploitation (fishing mortality F/FMSY,
where MSY = maximum sustainable yield) on the y- axis and stock size (biomass B/BMSY) on the x- axis in the final year (2021) for the range
of discard mortality rates assessed (0.0– 0.5); (C) effect of discard mortality rate on estimated intrinsic population growth rate(r); and (D)
effect of discard mortality rate on estimated carrying capacity (k).
Ye ar
All fishery removals (1000 metric tons)
1950 1957 1964 1971 1978 1985 1992 1999 2006 2013 2020
0
2
4
6
8
10
12 Discard mortality
0.5
0.4
0.3
0.2
0.1
0.0
(A)
Stock size
Exploitation
0.80.9 1.01.1 1.2
0.8
0.9
1.0
1.1
1.2
Discard mortality
0.5
0.4
0.3
0.2
0.1
0.0
(B)
Discard mortality
Intrinsic population growth rate (r)
0.00.1 0.20.3 0.40.5
0.40
0.45
0.50
0.55
0.60 (C)
Discard mortality
Carrying capacity (k)
0.00.1 0.20.3 0.
40
.5
10
20
30
40
50
60 (D)
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14 of 22
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GERVASI et al.
beginning as early as 1985, with very low catch rates ob-
served since the early 2000s (Gervasi et al.2022b). Our
stock assessment results align with the timing of this
observation and further highlight the need to develop a
management plan for this important fishery.
Crevalle Jack stock status and trends
Our base CMSY– BSM model revealed that the catch of
Florida Crevalle Jack has been at or above MSY almost
every year since 1989, with several years of overfishing
occurring, and that the stock has been in an overfished
state during almost every year since 2003. Stock size has
been gradually declining over time, while recreational
fishing effort appears to be continually increasing. Com-
mercial landings were relatively low throughout the time
series compared to recreational landings, and commercial
landings dropped considerably in the mid- 1990s (coinci-
dent with the commercial gill- net ban in Florida; Smith
et al.2003). The increasing recreational fishing effort is
somewhat surprising, as fishing guides reported that the
Crevalle Jack fishery in the Florida Keys is largely op-
portunistic and catch and release (Gervasi et al. 2022b).
However, in the statewide MRIP data, recreational anglers
report which species were primarily targeted on each fish-
ing trip; out of all Florida trips, Crevalle Jack were reported
as the 46th most targeted species out of 318 species listed
as primary targets. Therefore, the Crevalle Jack is in the top
15% of recreationally targeted species throughout the state.
Studies have shown that recreational landings ex-
ceed commercial landings for many fisheries (Coleman
et al.2004; Radford et al.2018; Lewin et al.2019; Shertzer
et al.2019), and there is growing evidence that recreational
fisheries can be responsible for declines in fish populations
and can have other biological impacts (Lewin et al.2006;
Brownscombe et al. 2019). Even in predominantly catch-
and- release fisheries, postrelease mortality and sublethal
effects on physiology can have substantial impacts on fish
populations (Rudershausen et al.2007; Cooke et al.2013).
Worldwide, the number of recreational anglers (Kear-
ney2002; Pawson et al.2008), the magnitude of recreational
catches (Coleman et al.2004; Felizola Freire et al.2020), and
the economic impact of recreational fishing (Arlinghaus
et al.2019) are increasing. Although recreational fisheries
provide funding for conservation efforts and connect soci-
ety with nature, thereby increasing public awareness and
appreciation of conservation concerns (Griffiths et al.2017;
Arlinghaus et al. 2019; Brownscombe et al. 2019), these
fisheries are prone to high uncertainty, which undermines
sustainable management (Shertzer et al.2019). Appropri-
ate management action that balances the social and ecolog-
ical dimensions of these fisheries is therefore vital.
In addition to increased fishing effort, other factors
may have contributed to the decline and may continue
to impact Crevalle Jack populations in the future. During
LEK interviews, fishing guides were asked to speculate on
potential reasons for the perceived decline in Crevalle Jack
catches, and loss of prey was the most commonly men-
tioned reason (followed by recreational harvest; Gervasi
et al.2022b). Poor water quality, increased predators, and
warmer winters were also potential factors mentioned by
multiple guides. Research has shown that regional climate
variability can lead to changes in the distribution and pro-
ductivity of fish species (Brander2007; Lotze et al.2019).
It is therefore possible that climate- induced shifts in prey
or predator species have contributed to shifts in Crevalle
Jack distributions in the region. Ecosystem- based man-
agement efforts in the South Atlantic and Gulf of Mexico
could contribute to more holistic management of species
such as the Crevalle Jack in the future (e.g., fisheries eco-
system plans; Levin et al.2018).
FIGURE Results of the base model in comparison with sensitivity analyses 2– 7 (S2– S7): (A) development of predicted relative total
biomass (B/BMSY, where MSY = maximum sustainable yield) for each model run; and (B) time series of exploitation (fishing mortality
F/FMSY) on the y- axis and stock size (B/BMSY) on the x- axis in the final year (2021) for each model run.
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
The timing and trajectory of Crevalle Jack exploitation
match the observations of recreational fishing guides in
Florida, some of whom began noticing a decline in Cre-
valle Jack catch rates as early as 1985 (Gervasi et al.2022b).
Most guides, however, noticed the decline in the early to
mid- 2000s, which corresponds to the year when stock
size began dipping below BMSY (2003). Additionally,
guides reported that the decline had been gradual, which
again matches the model results (i.e., even for the anal-
ysis using uninformed priors, stock size declined gradu-
ally from 1970 to 2021). This agreement between fishing
guide observations and model results provides confi-
dence in the stock assessment and highlights the benefits
of incorporating LEK into fisheries research. Consistency
between LEK and other data sources has been observed
in many studies (e.g., Poizat and Baran1997; Aswani and
Hamilton2004; Zukowski et al.2011; Rehage et al.2019;
Santos et al.2019; Bourdouxhe et al.2020), and the use of
LEK in fisheries research and management has increased
substantially over the years (Beaudreau and Levin2014).
A recent study by Shephard et al.(2021) showed that an-
gler LEK matched stock assessment results for four rec-
reational fisheries in Ireland, further demonstrating that
LEK can provide valuable, robust information about fish-
eries stock status and trends.
Importantly, research and management efforts that
rely on stakeholder input and collaboration are most suc-
cessful in situations of mutual trust and respect, which
can be difficult to build and maintain (Thornton and
Scheer2012). In our study, we solicited the aid of experi-
enced recreational fishing guides to fill in knowledge gaps
about the Crevalle Jack fishery and to help inform model
priors. Studies have shown that fishing guides are ideal
research partners, as they have substantial on- the- water
experience and a vested interest in fisheries conservation
(Kroloff et al.2019; Adkins2020; Gervasi et al.2022a). To
ensure continued trust and collaboration, fishing guides
were informed of the results of this study and its potential
management applications. As demonstrated by Gervasi
et al.(2022a), it is important to involve anglers through-
out the scientific research process and beyond to maintain
trusted partnerships. For cases in which there is general
distrust of science and management by key stakeholders,
efforts to build trust and maintain relationships are vital
to conducting LEK research and fisheries co- management
(Thornton and Scheer2012; Rubert- Nason et al.2021).
Sensitivity analyses
Fisheries management is commonly based on setting
target quotas or catch limits based on fisheries reference
points from stock assessments (Newman et al. 2015).
Uncertainty in model parameters that greatly affect the
estimation of reference points can lead to target setting
based on inaccurate estimations of stock status, thus
increasing the risk for either overfishing or underutiliz-
ing the resource (Dankel et al.2012; Cadrin et al. 2015;
Privitera- Johnson and Punt 2020). Sensitivity analysis is
a common approach used by stock assessment scientists
to understand aspects of model uncertainty (Privitera-
Johnson and Punt2020). Compared to our initial CMSY–
BSM model, none of the sensitivity analyses dramatically
altered the overall pattern of exploitation and stock size
over time or the estimated current stock status. In all mod-
els, exploitation increased over time, with harvest increas-
ing to levels at or above MSY at some point during the
time series. Stock size also generally decreased over time,
with overfishing occurring in all models, although the
number of years for which the stock was in an overfished
state varied depending on the model. Exploitation in 2021
was high for all models, with F2021/FMSY ranging from 0.93
to 1.81 (Table3). All models also showed that the stock
in 2021 was overfished (B2021/BMSY < 1). This model con-
sistency reveals high model precision and provides some
additional confidence in our stock assessment results.
However, there still may be unaccounted- for sources
of uncertainty (i.e., “unknown unknowns”; Drouineau
et al.2016) that could affect model accuracy.
Despite the consistency in overall trends among model
runs, estimated management reference points deviated
from the initial model for some of the sensitivity analyses.
Changing the discard mortality rate for our first analysis
had the greatest effect on reference points, and k increased
dramatically with an increase in the discard mortality rate.
This change in k led to a substantial impact on estimated
MSY and BMSY, which are important values needed to de-
termine fishery quotas. This analysis highlights the impor-
tance of estimating an accurate discard mortality rate for
fisheries that are predominantly catch and release. When
angling effort is high, catch- and- release fishing is often ap-
plied as a management solution for reducing angling im-
pacts on important fisheries (Cooke and Schramm2007).
Although catch- and- release fishing can provide many
benefits to fisheries when used appropriately (Arling-
haus et al. 2002, 2007), it can also have unintended and
unaccounted- for consequences (Cooke et al.2002; Cooke
and Suski2005). Several studies have shown that angling
can have a multitude of physiological effects on fish, re-
sulting in morbidity and mortality after release (Cooke
et al. 2002; Campbell et al.2010), and can increase vul-
nerability to predation (Holder et al.2020). Accurately ac-
counting for discard mortality in assessments of largely
catch- and- release fisheries is therefore vital.
Of the remaining sensitivity analyses, using unin-
formed priors had the greatest effect on estimates of k
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16 of 22
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GERVASI et al.
and BMSY, resulting in a much more optimistic view of
available biomass and stock status. According to this
version of the model, the stock was marginally over-
fished but not undergoing overfishing in 2021. Failure
to provide informed priors could therefore prevent man-
agement action from being taken, potentially leading to
continued overfishing and even stock collapse. Previous
research has shown that Bayesian methods (e.g., BSM)
are highly sensitive to misspecified priors and that well-
thought- out informative priors can considerably reduce
uncertainty (Punt and Hilborn 1997). Expert anglers
have been shown to provide accurate estimates of bio-
mass trends in many studies (Beaudreau and Levin2014;
Shephard et al.2021) and thus serve as a useful resource
for developing informative priors. In fact, research
has shown that synthesizing expert knowledge can be
the most powerful approach for selecting informative
model priors (Punt and Hilborn1997). Previous studies
the specifically employed the CMSY– BSM method have
used expert knowledge to inform the relative biomass
priors required by the model (Demirel et al.2020). The
results of this sensitivity analysis highlight the impor-
tance of the LEK component of our assessment frame-
work (Figure1).
Incorporating effort creep into the model also had
a significant impact on estimated reference points and
stock status. When accounting for a 2% linear increase
in catchability, the model resulted in a much more pes-
simistic view of exploitation and stock status, with B2021
being critically low. Effort creep (also called “technology
creep”) has been shown to significantly alter how fishing
impacts fish stocks (Marchal et al.2007; Scherrer and Gal-
braith2020), but creep factors are typically only estimated
to correct for the introduction of new technologies over
short periods of time. Therefore, applying a blanket effort
creep value to a long- term analysis is not ideal (Palomares
and Pauly2019). Unfortunately, there is a general lack of
quantitative data on the speed and magnitude with which
fishing power changes over time (Engelhard2016). Future
efforts to explicitly quantify changes in catchability due to
advancements in fishing technology could greatly improve
stock assessment models and inform better management.
Finally, excluding high- PSE data points from the MRIP
data also led to a more pessimistic stock status than the
base model due to the anomalously high MRIP catch esti-
mate for 2009. If this was a true spike in catch reflective of
a spike in abundance for that year, its cause is unknown.
A strong recruitment event was a possible cause. It is well
known that variability in juvenile recruitment rates due to
environmental variability can lead to substantial tempo-
ral heterogeneity in population abundance (Shelton and
Mangel2011). However, fishing guides did not mention
any particular spike in Crevalle Jack abundance in 2009 or
anything else that would explain the spike. Additionally,
the PSE of the MRIP estimate was above 50%, meaning
that the estimate was very imprecise. It is therefore more
likely that the catch estimate was based on a small sam-
ple size and is not a “true” reflection of total catch in that
year. Studies have shown that data quantity significantly
impacts stock assessment results (Chen et al.2003). This
sensitivity analysis further highlighted the importance of
considering sample size and the precision of catch esti-
mates when fisheries- dependent data are used to inform
stock assessment.
Implications for management
Our sensitivity analyses revealed some uncertainty in the
extent of overfishing that has occurred since 1950, but all
models showed stock size trending in a negative direction,
suggesting that management action is needed to halt the
decline in stock size. The current exploitation rate is also
at or slightly above MSY. Since the Crevalle Jack is cur-
rently an unregulated species in Florida (FWC2021) and
given that recreational fishing in the region is continually
increasing (Hanson and Sauls2011; Shertzer et al.2019),
it is likely that exploitation rates will continue to increase
to unsustainable levels if the fishery remains unregulated.
Importantly, with recreational fisheries the goal is not
always to maximize yield. Fishing guides in the Florida
Keys have observed that catch rates of Crevalle Jack have
declined below a desirable level in recent years (Gervasi
et al.2022b). Thus, although F2021 was just above FMSY and
B2021 was only slightly below BMSY for most of our model
runs, management regulations that bring catch rates back
up to desirable levels may be more beneficial to the guided
fishery as an industry than managing for MSY. Further
discussions with anglers as to what constitutes a desirable
level of catch will help managers to set appropriate refer-
ence points. Because the Crevalle Jack is an unregulated
species (i.e., there are no species- specific restrictions on
harvest) in all U.S. Gulf and Atlantic states within the spe-
cies' range, additional research into Crevalle Jack stock
structure and stock status in other areas is also a critical
next step.
Our suggested next steps for management include en-
gaging in cooperative research and co- management (Ka-
plan and McCay2004; Johnson and Van Densen2007) and
setting regulations on the Crevalle Jack recreational fish-
ery that are acceptable to the stakeholders and that follow
a precautionary approach. Beyond such steps, additional
research can aid in reducing uncertainty and providing
more concrete management recommendations. The results
of our sensitivity analyses revealed the importance of es-
timating an accurate discard mortality rate since the vast
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ASSESSMENT OF AN UNREGULATED FISHERY USING DATA-LIMITED TOOLS
majority of Crevalle Jack captured by recreational anglers
in Florida are released. Tagging studies that assess how
factors such as handling time, hooking location, depth,
and predator abundance influence postrelease survival
will aid in obtaining a better estimate of the survival rate
(Jiang et al.2007; Rudershausen et al.2007; Flaherty- Walia
et al.2016). Accounting for effort creep was also shown to
be incredibly important. Therefore, getting a better handle
on how fishing technology and subsequent catchability of
Crevalle Jack may have changed over time should be an-
other research goal. This could potentially be accomplished
via angler interviews and/or analysis of trends in the adop-
tion and use of new fishing technologies in the region
(e.g., Marchal et al.2007). Finally, accurate delineation of
stock boundaries is an important part of stock assessment
(Ying et al.2011; Berger et al. 2021). Preliminary acous-
tic telemetry research in Florida has revealed that Crevalle
Jack make regular long- range movements throughout the
state and that some individuals even cross state boundaries
into other states within the Gulf of Mexico (C. L. Gervasi,
unpublished data). These results suggest that to encom-
pass the entire stock, the catch and abundance time series
may need to be expanded to include data from other states.
However, according to the MRIP data, approximately 95%
of the Crevalle Jack captured by recreational anglers in
Gulf of Mexico and South Atlantic waters are captured
in Florida. Thus, even if individual fish migrate between
state boundaries, fishing operations in other states are less
likely to impact stock status since the majority of the Cre-
valle Jack fishery operates in Florida. For this reason, a
stock unit extending beyond the state of Florida was not
considered for the assessment conducted herein. However,
as the acoustic telemetry data continue to reveal patterns of
Crevalle Jack movements and migrations, the CMSYBSM
model could be re- run if necessary to account for changes
in estimated stock boundaries. As new data about the spe-
cies and the fishery are collected, the FishPath assessment
questionnaire can also be updated and other data- limited
assessment methods can be explored and compared. The
three- prong assessment approach outlined herein first uses
the FishPath tool to select an assessment method, then
conducts an assessment using the chosen method, and fi-
nally runs sensitivity analyses for unknown or uncertain
parameters. Local ecological knowledge permeates each
step, rapidly filling in knowledge gaps that would other-
wise take years of additional research and data collection
to fill. This approach can easily be included as part of an
adaptive management plan and can be applied to other un-
regulated species and in other regions.
ACKNOWLEDGMENTS
We are grateful to the recreational fishing guides who
contributed their experiences and expertise to this study
as well as the Lower Keys Guides Association and Florida
Keys Fishing Guides Association for assistance in identi-
fying key informants. We especially thank C. Bradshaw
for information about the Crevalle Jack commercial
fishery and the two anonymous reviewers for comments
that improved the manuscript. Much appreciation goes
to the Nature Conservancy and the creators and devel-
opers of the FishPath tool. We also appreciate R. Froese,
N. Demirel, G. Coro, and H. Winker for developing the
CMSY+ and BSM models and for their thorough docu-
mentation of the method and reproducible code. This
study was developed in collaboration with the Florida
Coastal Everglades Long- Term Ecological Research
Program under National Science Foundation Grant No.
DEB- 1832229, and this work was reviewed and deemed
exempt by the FIU Institutional Review Board (Proto-
col Exemption IRB- 19- 0157; May 7, 2019). This material
is based upon work supported by the National Science
Foundation under Grant No. HRD- 1547798 and Grant
No. HRD- 2111661. These NSF Grants were awarded to
Florida International University as part of the Centers of
Research Excellence in Science and Technology (CREST)
Program. Funding was provided by the Lower Keys
Guides Association, private donation, the Everglades
Foundation FIU ForEverglades Graduate Scholarship,
and the FIU Graduate School Dissertation Year Fellow-
ship. This is Contribution Number 1609 from the Insti-
tute of Environment, a Preeminent Program at Florida
International University.
CONFLICT OF INTEREST STATEMENT
There is no conflict of interest declared in this article.
DATA AVAILABILITY STATEMENT
The data underlying this article will be shared upon rea-
sonable request to the corresponding author.
ETHICS STATEMENT
All ethical guidelines were followed and no animals were
handled in the development of this study.
ORCID
Carissa L. Gervasi https://orcid.
org/0000-0003-1590-9332
Mandy Karnauskas https://orcid.
org/0000-0002-6631-0592
Adyan Rios https://orcid.org/0000-0003-0112-3080
Rolando O. Santos https://orcid.
org/0000-0002-3885-9406
W. Ryan James https://orcid.org/0000-0002-4829-7742
Ryan J. Rezek https://orcid.org/0000-0002-3054-6384
Jennifer S. Rehage https://orcid.
org/0000-0003-0009-6906
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